Discriminating alternative stock–recruitment models and evaluating uncertainty in model structure

Fish stock–recruitment (S–R) assessment is one of the most essential keystones for fisheries management. Yet the analysis involves a variety of uncertainties. Amidst these difficulties, uncertainty in model structure is perhaps the most problematical to investigate because no rigorous statistical techniques can be used to explore the fundamental biological processes in S–R relationships. In this paper, I used computer simulations to investigate: (1) the differences between the estimated parameters of alternative S–R models as a function of stock characteristics: population growth rate, data range, fishing mortality, and process noise; and (2) the probability of selecting a correct model using information criteria. Two popular S–R functions, the Ricker and the Beverton–Holt models, were used as examples. Time series data were generated from a known S–R model and fitted by alternative models. The results show that when the two models fit the data similarly well, significant differences in parameters existed between the alternative models. The Ricker model tended to underestimate the population growth rate (initial slope) and the carrying capacity parameter, whereas the Beverton–Holt model overestimated these parameters. The management quantities (e.g., optimal virgin stock size) produced by one model were more conservative (i.e., larger optimal stock size or lower optimal harvest rate) under some conditions but became less conservative under other conditions. The differences between the alternative models were functions of the population growth rate, long-term fishing mortality, and data range of the stock size. The correct and incorrect models were statistically indistinguishable. For typical fishery data the probability of selecting the correct model based on information criteria was approximately 0.70 for the Ricker model and 0.61 for the Beverton–Holt model.     

Population reproductive potential: Evaluation of long-term stock productivity

In order to avoid recruitment overfishing, fish stocks must have sufficient reproductive ability. The spawning stock biomass (SSB), which ignores the value of immature fish, is widely used as an index of stock sustainability. From the perspective of sustainability, immediate reproduction, as well as future spawning, must be considered. We developed an index of long-term stock productivity, called the population reproductive potential (PRP). PRP is defined as the expected total reproductive value of the standing stock. We used PRP to assess the western Atlantic bluefin tuna (WBT) stock. The trends in SSB, numbers (N), biomass and PRP of WBT are inconsistent when compared to each other, due to fluctuation in age composition. We evaluated the long-term productivity of WBT by computer simulation and compared the result with trends in the abundance indices. The result of the computer simulation was highly consistent with the trend in the PRP. Short-term trends in SSB and N often do not reflect long-term stock trends, because they are highly sensitive to age-composition dynamics. The PRP is useful for evaluating stock trends, especially when the age composition is unstable.     

象山港黄姑鱼增殖放流效果评估及增殖群体利用方式优化

以回捕渔获重量和对繁殖群体的补充能力为评价指标,借助标志放流-回捕实验,利用模型模拟分析方法,定量评估浙江象山港黄姑鱼(Nibea albiflora)的增殖放流效果;同时,结合其增殖目标定位,探索构建生态高效的增殖群体利用方式。研究结果表明,象山港黄姑鱼增殖群体的捕捞死亡系数为1.31,在该捕捞强度下,11055尾增殖放流鱼苗可产生737 kg回捕渔获收益,同时还向增殖水域补充了554尾初次性成熟个体,增殖放流活动在一定程度上起到了修复象山港黄姑鱼资源、促进渔民增产增收的效果。结果表明,捕捞强度过大是制约象山港黄姑鱼增殖放流功效发挥的重要因素,捕捞死亡系数应降至0.46,同批次黄姑鱼增殖放流所能提供的回捕渔获重量和性成熟个体尾数可分别较现行捕捞强度提升41.49%和326.90%。综上述,象山港黄姑鱼增殖放流的生态和经济功效较为显著,合理降低对增殖群体的捕捞强度是进一步提升其增殖放流效果的必要措施。     

Does the Ricker or Beverton and Holt type of stock-recruitment relationship truly exist?

ABSTRACT:   This study investigated the main causes of population abundance fluctuations. Particularly, attention was paid to whether a density-dependant factor, such as a stock-recruitment relationship (S-R relationship) or a density-independent factor such as an environmental factor, is more important. Using data pertaining to the number of eggs of the Pacific stock mackerel and information about regime shifts and sea surface temperature, the shape of the S-R relationship was discussed and these shapes with the results of simulation trails were compared. Further other historical S-R relationship data were analyzed. The results are as follows: (i) a new mechanism that causes population fluctuations could be proposed, that is, (a) the recruitment is proportional to the spawning stock biomass (SSB) and the relation is expressed by several lines with the same slope, and (b) the shift between the lines occurs due to environmental conditions; and (ii) the density-dependent S-R relationship, which suggests that recruitment decreased due to high density in SSB, proposed by Ricker or Beverton and Holt, would not exist.     

Stock–environment–recruitment models for North Atlantic albacore (Thunnus alalunga)

Different stock–recruitment models were fitted to North Atlantic albacore (Thunnus alalunga) recruitment and spawning stock biomass data. A classical density dependence hypothesis, a recent environmental‐dependence hypothesis and a combination of both were considered. For the latter case, four stock–environment–recruitment models were used: Ricker, Beverton‐Holt, Deriso's General Model (modified to take into account environmental effects) and conditioned Neural Networks. Cross‐validation analysis showed that the modified Deriso model had the best predictive capability. It detected an inverse effect of the North Atlantic Oscillation (NAO) on recruitment, a Ricker‐type behaviour with density dependent overcompensation when environmental conditions are unfavourable and a Beverton–Holt‐type behaviour towards an asymptotic recruitment carrying capacity with favourable environmental conditions. The Neural Network model also detected that under favourable environmental conditions high spawning stock biomass does not necessarily have a depensatory effect on recruitment. Moreover, they suggest that under extremely favourable environmental conditions, albacore recruitment could increase well above the asymptotic carrying capacity predicted by Beverton–Holt‐type models. However, the general decrease in spawning stock biomass in recent years and increasing NAO trends suggest that there is low probability of exceptionally large recruitment in the future and instead there is a danger of recruitment overfishing.     

珠江禁渔对广东鲂资源补充群体的影响分析

继我国各大水系实施禁渔期制度之后,珠江水系于2011年4月1日至6月1日实现首次禁渔。广东鲂是珠江中下游地区主要的经济鱼类之一,为分析珠江实施禁渔期制度对广东鲂资源补充群体的影响,于珠江实施禁渔制度前(2006—2010年)和禁渔制度后(2011—2012年)在珠江下游肇庆江段设立固定采样点,利用定量弶网对流经该江段的广东鲂鱼苗补充群体进行调查。结果显示,禁渔前,珠江广东鲂鱼苗集中出现在4月中下旬至10月中旬,持续时间为(189±11)d,高峰期为6—8月。禁渔期制度实施之后,全年广东鲂鱼苗出现的起始时间变化不大,但是结束时间提前,持续时间略有缩短,4—5月份广东鲂鱼苗密度明显增大。广东鲂鱼苗总量及其在鱼苗补充群体中的比例均有所提高。根据禁渔前广东鲂鱼苗发生量与径流量之间存在的回归关系:LgY=0.988Lnx-4.932(R2=0.365,P0.01),结合禁渔后实际调查结果,对禁渔的实际作用进行了估算,2011年和2012年由于禁渔分别增加了9.43×108和651.81×108尾广东鲂鱼苗。研究表明,在径流量有保障的前提下,目前的禁渔制度可以有效增加广东鲂鱼苗补充群体资源量。     

Causes of walleye pollock (Theragra chalcogramma) recruitment decline in the northern Sea of Japan: implications for stock management

Recruitment of the northern Japan Sea stock (JSS) of walleye pollock has been decreasing since around 1990. In this study, I analyzed the factors causing this decrease in recruitment by investigating the relationship between recruitment, spawning stock biomass (SSB) and environmental factors using a generalized additive model (GAM). GAM fit to the data showed the importance of SSB, sea surface temperature (SST), ocean current strength (Tsushima Warm Current) and wind intensity (Asian monsoon) in determining the recruitment. Of these, the relationship between SSB and recruitment was positive and not negatively density‐dependent. On the other hand, the recruitment was negatively related to SST and ocean current strength, and a dome‐shaped relationship was observed between wind intensity and recruitment. Since around 1990, the values of SST and ocean current strength have mostly been high and that of wind intensity mostly low. In addition, SSB has been decreasing since the late 1990s. It is likely that the recruitment decline of JSS after approximately 1990 has been caused by warm water temperature, strong Tsushima Warm Current and weak Asian monsoon, and that the recent decrease in SSB has amplified this recruitment decline. According to the model’s estimation, a recruitment recovery due to environmental improvement will be highly restricted as long as SSB remains at its current low level. Significant recovery of SSB is urgently needed for JSS.     

Effects of biological,economic and management factors on tuna and billfish stock status

Commercial tunas and billfishes (swordfish, marlins and sailfish) provide considerable catches and income in both developed and developing countries. These stocks vary in status from lightly exploited to rebuilding to severely depleted. Previous studies suggested that this variability could result from differences in life‐history characteristics and economic incentives, but differences in exploitation histories and management measures also have a strong effect on current stock status. Although the status (biomass and fishing mortality rate) of major tuna and billfish stocks is well documented, the effect of these diverse factors on current stock status and the effect of management measures in rebuilding stocks have not been analysed at the global level. Here, we show that, particularly for tunas, stocks were more depleted if they had high commercial value, were long‐lived species, had small pre‐fishing biomass and were subject to intense fishing pressure for a long time. In addition, implementing and enforcing total allowable catches (TACs) had the strongest positive influence on rebuilding overfished tuna and billfish stocks. Other control rules such as minimum size regulations or seasonal closures were also important in reducing fishing pressure, but stocks under TAC implementations showed the fastest increase of biomass. Lessons learned from this study can be applied in managing large industrial fisheries around the world. In particular, tuna regional fisheries management organizations should consider the relative effectiveness of management measures observed in this study for rebuilding depleted large pelagic stocks.     

Is spawning stock biomass a robust proxy for reproductive potential?

Reference points based on fishing mortality (F) and spawning stock biomass (SSB) are a requirement of many fisheries management frameworks. SSB is assumed to be a proxy for stock reproductive potential (SRP). Limit reference points based on SSB are used to indicate the level of biomass below which productivity is affected. SSB fails to account for changes in fecundity, egg viability and sex ratio, and it has been argued that total egg production (TEP) provides a better reflection of SRP. We explore how accounting for TEP impacts limit reference points and evidence for a relationship between stock and recruit. Time series of SSB and TEP are compared for three North Sea stocks: cod (Gadus morhua), herring (Clupea harengus) and plaice (Pleuronectes platessa). Dynamics based on TEP are different from those based on SSB for cod and plaice, but the stock–recruit relationships were not ‘improved’ using TEP. Shifts in productivity (spawner per recruit) occur in all three time series and SSB underestimated uncertainty. Yet again, it was shown that assumptions of stationarity about fish population productivity are incorrect. We argue that the use of TEP does improve the realism in our understanding of stock dynamics, and demographically, more complex management strategy evaluation is required to develop management procedures that are robust to uncertainty and integrate F and the demographic health of a stock. Empirical feedback control systems based on fisheries independent indices including surveys of eggs, larvae, recruits, juveniles or spawning adults should be evaluated and compared to traditional approaches.     

Reliability of stock size estimates from adaptive framework virtual population analysis

ABSTRACT:   The stock size estimates from adaptive framework virtual population analysis are sometimes very sensitive to the model's assumptions. The uniqueness and precision of stock size estimates from adaptive framework virtual population analysis are examined analytically and numerically. The results indicate that (i) when the fishing mortalities remain constant for all years, the stock size estimates are not determined uniquely without additional assumptions, and (ii) when the cumulative fishing mortalities are similar among cohorts, the precision of stock size estimates will be poor, even if the abundance indices are precise and informative. For the precise estimation of stock size, it is necessary that fishing mortalities vary among years, that cumulative fishing mortalities vary among cohorts and that auxiliary information such as selectivity is available in addition to precise abundance indices.     

适用于数据缺乏渔业的资源评估方法研究进展

开展渔业资源评估研究是制定渔业可持续发展策略的重要前提,而数据有限是全球渔业资源评估面临的普遍挑战.传统资源评估方法具有数据需求量大、要求高等特点,无法应用于数据缺乏渔业的资源评估中.数据缺乏方法(data-limited method)可结合少量易获得数据和相关历史生物学信息对渔业资源状况、生物学参考点以及资源量等进...     

Effects of process and/or observation errors on the stock–recruitment curve and the validity of the proportional model as a stock–recruitment relationship

We examined the effects of process and observation errors on the selection of the stock–recruitment relationship (SRR) curve using simulations. When the process and observation errors were added to both spawning stock biomass and recruitment, the results were as follows: (1) When the proportional model was set as the true SRR model, there was a high probability that the Ricker or Beverton and Holt model was selected in response to the errors; (2) When the Ricker or Beverton and Holt model was set as the true SRR model, the proportional model was seldom selected in response to the errors; (3) The proportional SRR model should be accepted as the optimum SRR model for the Pacific stock of Japanese sardine; (4) We should use an SRR model that is constructed from more than two independent variables (spawning stock biomass, environmental factors, etc.) when we discuss management of fisheries resources.     

基于SPiCT模型对数据有限短生命周期阿根廷滑柔鱼的资源评估

以西南大西洋阿根廷滑柔鱼(Illex argentinus)为研究对象,基于连续时间的随机剩余产量模型(a stochastic surplus production model in continuous time,SPiCT),分析了6种方案下参数估计的变化及其对资源评估的影响。对比6种方案中阿根廷滑柔鱼的产量和单位捕捞努力渔获量(catch per unit effort,CPUE)的估计值与观测值间的最小残差平方和,方案3(设置了K、r、q的先验分布)为最适方案。相应的资源评估结果显示,2010年西南大西洋阿根廷滑柔鱼捕捞死亡系数小于最大持续产量时捕捞死亡系数F_(MSY),渔获量小于最大持续产量MSY,预期平衡生物量EEB大于最大持续产量时的生物量B_(MSY),这表明该资源在2010年尚未被过度开发利用。SPiCT模型综合考虑了环境因子、种群间相互作用和网具选择性等因素引起的观测和过程误差,较S、F-EDSP、S-F-EDSP模型及其他离散模型对数据要求低,计算方法简单,更适合数据有限、短生命周期渔业种类的资源评估。另外,可捕系数q值的设置严重影响了SPiCT模型K、B的估计,优化估计可捕系数q将有利于提高其资源评估的准确性。     

Are stock assessment methods too complicated?

This critical review argues that several methods for the estimation and prediction of numbers‐at‐age, fishing mortality coefficients F, and recruitment for a stock of fish are too hard to explain to customers (the fishing industry, managers, etc.) and do not pay enough attention to weaknesses in the supporting data, assumptions and theory. The review is linked to North Sea demersal stocks. First, weaknesses in the various types of data used in North Sea assessments are summarized, i.e. total landings, discards, commercial and research vessel abundance indices, age‐length keys and natural mortality (M). A list of features that an ideal assessment should have is put forward as a basis for comparing different methods. The importance of independence and weighting when combining different types of data in an assessment is stressed. Assessment methods considered are Virtual Population Analysis, ad hoc tuning, extended survivors analysis (XSA), year‐class curves, catch‐at‐age modelling, and state‐space models fitted by Kalman filter or Bayesian methods. Year‐class curves (not to be confused with ‘catch‐curves’) are the favoured method because of their applicability to data sets separately, their visual appeal, simple statistical basis, minimal assumptions, the availability of confidence limits, and the ease with which estimates can be combined from different data sets after separate analyses. They do not estimate absolute stock numbers or F but neither do other methods unless M is accurately known, as is seldom true.     

Forecasting fish stock dynamics under climate change: Baltic herring (Clupea harengus) as a case study

Climate change and anthropogenic disturbances may affect marine populations and ecosystems through multiple pathways. In this study we present a framework in which we integrate existing models and knowledge on basic regulatory processes to investigate the potential impact of future scenarios of fisheries exploitation and climate change on the temporal dynamics of the central Baltic herring stock. Alternative scenarios of increasing sea surface temperature and decreasing salinity of the Baltic Sea from a global climate model were combined with two alternative fishing scenarios, and their direct and ecosystem‐mediated effects (i.e., through predation by cod and competition with sprat) on the herring population were evaluated for the period 2010–2050. Gradual increase in temperature has a positive impact on the long‐term productivity of the herring stock, but it has the potential to enhance the recovery of the herring stock only in combination with sustainable fisheries management (i.e., F_msy). Conversely, projections of herring spawning stock biomass (SSB) were generally low under elevated fishing mortality levels (F_high), comparable with those experienced by the stock during the 1990s. Under the combined effects of long‐term warming and high fishing mortality uncertainty in herring SSB projections was higher and increasing for the duration of the forecasts, suggesting a synergistic effect of fishery exploitation and climate forcing on fish populations dynamics. Our study shows that simulations of long‐term fish dynamics can be an informative tool to derive expectations of the potential long‐term impact of alternative future scenarios of exploitation and climate change.     

东海北部哈氏仿对虾资源量评估

根据2005～2008年东海北部哈氏仿对虾的生物学数据,应用FISATⅡ软件的ElEFANⅠ技术计算生长参数,得到VonBertalanffy生长方程的相关参数为L∞=123.4mm,K=1.70,t0=-0.16,自然死亡系数M=2.55,总死亡系数Z=5.81,捕捞死亡系数F=3.26。应用长度结构的实际种群分析,得出稳定状态下资源量为2.86×104吨。利用Beverton-Holt和Thompson-Bell模型对单位补充量的产量Y/R和单位补充量的产值Yv/R进行分析,表明在现行状态下,该群体未出现过度捕捞现象。综合考虑当前可能的捕捞强度及经济效益,在有利于保持较高产量的前提下,建议在适当降低捕捞强度的同时把哈氏仿对虾的开捕体长由55.6mm增加至74.0mm。     

Stock assessment of Japanese eels using Japanese abundance indices

The paper compiles a catch history of Japanese eels Anguilla japonica in East Asia and some Japanese relative abundance series. Maximum likelihood estimates of stock abundance of eels have been obtained using the abundance series and various biological parameters, such as growth, maturity and natural mortality. Age- and sex-structured models have been used to express the dynamics of stock abundance, and the Beverton and Holt model has been used to express the relationship between stock and recruitment. Data for estimations are standardized catch per unit effort of commercial fishery for exploitable stock (1954–2006 and 1968–2008) and for glass eel (1954–2010, 1972–2004, and 1973–1997). From the results of the base case scenario of estimations, the estimated stock size of individuals aged ≥1 year was 18.7 thousand tons in 2010, which was 24 % of the carrying capacity. The estimated stock size has recovered since 1990. Maximum sustainable yield was 4,180 tons if only the exploitable stock were utilized, and 266 tons if only the glass eel were utilized. These results and issues relating to estimation and management for reducing the fishery impact on stock are discussed.     

Potential yield estimates of unexploited pelagic fish stocks in Lake Malawi

The biomass of the offshore pelagic fish of Lake Malawi was estimated at 168 400 t and is currently unexploited by a fishery. This biomass comprises, by weight, 81% Cichlidae, 15% catfish and 4% Cyprinidae. A Beverton and Holt dynamic pool model was used to estimate potential sustainable yields and to forecast likely changes to the size and age structure of the fish populations under various levels of fishing effort. The total sustainable yield from the offshore fish population was estimated at 34 000 t yr^–1 and, if harvested, would almost double the yield of fish from the lake. Potential yields on an areal basis are low (14.2 kg ha^–1 yr^–1), but are consistent with expectations from studies of pelagic ecosystem productivity.     

Population trends of bycatch species reflect improving status of target species

Synthesis studies of fish stocks worldwide suggest improving status of mainly target species that are fully assessed. Other analyses, primarily based on catch data alone, but which include a wider range of species as well as bycatch, present a different view. Catch‐only analyses could be more robust if fishery‐independent data were used and discards accounted for. We develop a model that uses only survey biomass at length and landings data to estimate fishing mortality, spawning stock biomass (SSB) and discards. An analysis of species from the North Sea shows the model results compare well with most fully assessed stocks. When applied to bycatch species with limited data, trends in fishing mortality and SSB typically reflect those of the target species. In the last decade, mean fishing mortality rates have tended to decline, while mean SSB has increased. Despite increasing SSB, recent mean recruitment appears to have been lower than previously which may limit future biomass recovery. Species usually associated with more northerly distributions appear to show the greatest effect of weaker recruitment, which may be linked to climate. Estimated discards have tended to decline in magnitude as a result of reduced fishing mortality and associated lower total catches. The model offers a simple way to use both landings and survey data to obtain more detailed population trends for data limited species.     

渔业资源评估模型的研究现状与展望

制定合理的渔业管理计划, 实现渔业资源的可持续利用, 需对渔业资源进行科学的评估, 而渔业资源评估模型则是进行渔业资源评估的重要工具。随着计算机计算能力的提高及在多学科交叉的推动下, 近30多年来渔业资源评估模型得到了快速发展。随着渔业资源评估模型日益复杂、多样化, 模型的选择、使用难度也相应地增加了, 而模型的不恰当运用则可能导致渔业资源管理的失误。本文对渔业资源评估模型的基本结构、主要类型及参数估计所使用的统计模型等方面进行了全面回顾, 介绍了目前使用的主要渔业资源评估模型, 展示了渔业资源评估模型的发展历程及所取得的进展。同时, 文章对渔业资源评估模型中存在的问题进行了探讨, 并对其未来的发展进行了展望。